ARTICLE UPDATE - Unconscious fear influences emotional awareness of faces and voices

B. de Gelder, J. S. Morris and R. J. Dolan

PNAS, 102, 18682-18687

Nonconscious recognition of facial expressions opens an intriguing possibility that two emotions can be present together in one brain with unconsciously and consciously perceived inputs interacting. We investigated this interaction in three experiments by using a hemianope patient with residual nonconscious vision. During simultaneous presentation of facial expressions to the intact and the blind field, we measured interactions between conscious and nonconsciously recognized images. Fear-specific congruence effects were expressed as enhanced neuronal activity in fusiform gyrus, amygdala, and pulvinar. Nonconscious facial expressions also influenced processing of consciously recognized emotional voices. Emotional congruency between visual and an auditory input enhances activity in amygdala and superior colliculus for blind, relative to intact, field presentation of faces. Our findings indicate that recognition of fear is mandatory and independent of awareness. Most importantly, unconscious fear recognition remains robust even in the light of a concurrent incongruent happy facial expression or an emotional voice of which the observer is aware.

ARTICLE UPDATE - Neural mechanism for judging the appropriateness of facial affect

Ji-Woong Kim, Jae-Jin Kim, Bum Seok Jeong, Seon Wan Ki, Dong-Mi Im, Soo Jung Lee and Hong Shick Lee

Cognitive Brain Research, 25, 659-667

Questions regarding the appropriateness of facial expressions in particular situations arise ubiquitously in everyday social interactions. To determine the appropriateness of facial affect, first of all, we should represent our own or the other's emotional state as induced by the social situation. Then, based on these representations, we should infer the possible affective response of the other person. In this study, we identified the brain mechanism mediating special types of social evaluative judgments of facial affect in which the internal reference is related to theory of mind (ToM) processing. Many previous ToM studies have used non-emotional stimuli, but, because so much valuable social information is conveyed through nonverbal emotional channels, this investigation used emotionally salient visual materials to tap ToM. Fourteen right-handed healthy subjects volunteered for our study. We used functional magnetic resonance imaging to examine brain activation during the judgmental task for the appropriateness of facial affects as opposed to gender matching tasks. We identified activation of a brain network, which includes both medial frontal cortex, left temporal pole, left inferior frontal gyrus, and left thalamus during the judgmental task for appropriateness of facial affect compared to the gender matching task. The results of this study suggest that the brain system involved in ToM plays a key role in judging the appropriateness of facial affect in an emotionally laden situation. In addition, our result supports that common neural substrates are involved in performing diverse kinds of ToM tasks irrespective of perceptual modalities and the emotional salience of test materials.

ARTICLE UPDATE - Doing the right thing: A common neural circuit for appropriate violent or compassionate behavior

John A. King, R. James R. Blair, Derek G.V. Mitchell, Raymond J. Dolan and Neil Burgess

NeuroImage, in press

Humans have a considerable facility to adapt their behavior in a manner that is appropriate to social or societal context. A failure of this ability can lead to social exclusion and is a feature of disorders such as psychopathy and disruptive behavior disorder. We investigated the neural basis of this ability using a customized video game played by 12 healthy participants in an fMRI scanner. Two conditions involved extreme examples of context-appropriate action: shooting an aggressive humanoid assailant or healing a passive wounded person. Two control conditions involved carefully matched stimuli paired with inappropriate actions: shooting the person or healing the assailant. Surprisingly, the same circuit, including the amygdala and ventromedial prefrontal cortex, was activated when participants acted in a context-appropriate manner, whether being compassionate towards an injured conspecific or aggressive towards a violent assailant. The findings indicate a common system that guides behavioral expression appropriate to social or societal context irrespective of its aggressive or compassionate nature.

ARTICLE UPDATE - How brains beware: neural mechanisms of emotional attention

Patrik Vuilleumier

Trends in Cognitive Sciences, in press

Emotional processes not only serve to record the value of sensory events, but also to elicit adaptive responses and modify perception. Recent research using functional brain imaging in human subjects has begun to reveal neural substrates by which sensory processing and attention can be modulated by the affective significance of stimuli. The amygdala plays a crucial role in providing both direct and indirect top-down signals on sensory pathways, which can influence the representation of emotional events, especially when related to threat. These modulatory effects implement specialized mechanisms of ‘emotional attention’ that might supplement but also compete with other sources of top-down control on perception. This work should help to elucidate the neural processes and temporal dynamics governing the integration of cognitive and affective influences in attention and behaviour.

ARTICLE UPDATE - Dissociable networks for the expectancy and perception of emotional stimuli in the human brain

Felix Bermpohl, Alvaro Pascual-Leone, Amir Amedi, Lotfi B. Merabet, Felipe Fregni, Nadine Gaab, David Alsop, Gottfried Schlaug and Georg Northoff

NeuroImage, in press

William James posited that comparable brain regions were implicated in the anticipation and perception of a stimulus; however, dissociable networks (at least in part) may also underlie these processes. Recent functional neuroimaging studies have addressed this issue by comparing brain systems associated with the expectancy and perception of visual, tactile, nociceptive, and reward stimuli. In the present fMRI study, we addressed this issue in the domain of pictorial emotional stimuli (IAPS). Our paradigm involved the experimental conditions emotional expectancy, neutral expectancy, emotional picture perception, and neutral picture perception. Specifically, the emotional expectancy cue was uncertain in that it did not provide additional information regarding the positive or negative valence of the subsequent picture. Neutral expectancy and neutral picture perception served as control conditions, allowing the identification of expectancy and perception effects specific for emotion processing. To avoid contamination of the perception conditions by the preceding expectancy periods, 50% of the pictorial stimuli were presented without preceding expectancy cues. We found that the emotional expectancy cue specifically produced activation in the supracallosal anterior cingulate, cingulate motor area, and parieto-occipital sulcus. These regions were not significantly activated by emotional picture perception which recruited a different neuronal network, including the amygdala, insula, medial and lateral prefrontal cortex, cerebellum, and occipitotemporal areas. This dissociation may reflect a distinction between anticipatory and perceptive components of emotional stimulus processing.

ARTICLE UPDATE - Changes in effective connectivity during incidental and intentional perception of fearful faces

G. de Marco, M. de Bonis, P. Vrignaud, M.C. Henry-Feugeas and I. Peretti

NeuroImage, in press

The present fMRI study examined effective connectivity within an emotional network composed of three brain areas: Amygdala (AMY), Anterior Cingulate Cortex (AAC) and Orbito-Frontal (OFC) in processing fearful faces. Two tasks: an incidental perception (gender identification) and an intentional detection (effortful discrimination) task were performed by 14 and 10 young healthy volunteers, respectively. Participants were scanned while viewing fearful, neutral and ambiguous facial expressions. Effective connectivity was assessed using Structural Equation Modeling (SEM). Results show that the hypothetical network fits the experimental data for both tasks and in both hemispheres. The comparison between Tasks 1 and 2 reveals significant differences in strength and direction of the connectivity patterns for the left and to a less stringent threshold for the right hemisphere. The path coefficients analysis suggests that the fearful information generated in AMY, reaches the OFC through the ACC in incidental perception, while in intentional perception, the route followed is in a reverse direction from OFC to ACC. Our findings confirm a differential brain connectivity between incidental and intentional processing of fearful faces.

ARTICLE UPDATE - “We All Look the Same to Me”: Positive Emotions Eliminate the Own-Race Bias in Face Recognition

Johnson, Kareem J., Fredrickson, Barbara L.

Psychological Science, 16, 875-881.

Extrapolating from the broaden-and-build theory, we hypothesized that positive emotion may reduce the own-race bias in facial recognition. In Experiments 1 and 2, Caucasian participants (N= 89) viewed Black and White faces for a recognition task. They viewed videos eliciting joy, fear, or neutrality before the learning (Experiment 1) or testing (Experiment 2) stages of the task. Results reliably supported the hypothesis. Relative to fear or a neutral state, joy experienced before either stage improved recognition of Black faces and significantly reduced the own-race bias. Discussion centers on possible mechanisms for this reduction of the own-race bias, including improvements in holistic processing and promotion of a common in-group identity due to positive emotions.

ARTICLE UPDATE - Attention modulates the processing of emotional expression triggered by foveal faces

Amanda Holmes, Monika Kiss and Martin Eimer

Neuroscience Letters, 394, 48-52.

To investigate whether the processing of emotional expression for faces presented within foveal vision is modulated by spatial attention, event-related potentials (ERPs) were recorded in response to stimulus arrays containing one fearful or neutral face at fixation, which was flanked by a pair of peripheral bilateral lines. When attention was focused on the central face, an enhanced positivity was elicited by fearful as compared to neutral faces. This effect started at 160 ms post-stimulus, and remained present for the remainder of the 700 ms analysis interval. When attention was directed away from the face towards the line pair, the initial phase of this emotional positivity remained present, but emotional expression effects beyond 220 ms post-stimulus were completely eliminated. These results demonstrate that when faces are presented foveally, the initial rapid stage of emotional expression processing is unaffected by attention. In contrast, attentional task instructions are effective in inhibiting later, more controlled stages of expression analysis.

ARTICLE UPDATE - Arousal and valence effects on event-related P3a and P3b during emotional categorization.

Delplanque S, Silvert L, Hot P, Rigoulot S, Sequeira H.

International Journal of Psychophysiology, in press

Due to the adaptive value of emotional situations, categorizing along the valence dimension may be supported by critical brain functions. The present study examined emotion-cognition relationships by focusing on the influence of an emotional categorization task on the cognitive processing induced by an oddball-like paradigm. Event-related potentials (ERPs) were recorded from subjects explicitly asked to categorize along the valence dimension (unpleasant, neutral or pleasant) deviant target pictures embedded in a train of standard stimuli. Late positivities evoked in response to the target pictures were decomposed into a P3a and a P3b and topographical differences were observed according to the valence content of the stimuli. P3a showed enhanced amplitudes at posterior sites in response to unpleasant pictures as compared to both neutral and pleasant pictures. This effect is interpreted as a negativity bias related to attentional processing. The P3b component was sensitive to the arousal value of the stimulation, with higher amplitudes at several posterior sites for both types of emotional pictures. Moreover, unpleasant pictures evoked smaller amplitudes than pleasant ones at fronto-central sites. Thus, the context updating process may be differentially modulated by the affective arousal and valence of the stimulus. The present study supports the assumption that, during an emotional categorization, the emotional content of the stimulus may modulate the reorientation of attention and the subsequent updating process in a specific way.

ARTICLE UPDATE - Contributions of the Amygdala to Emotion Processing: From Animal Models to Human Behavior

Phelps, E., & LeDoux, J.

Neuron, 48, 175-187

Research on the neural systems underlying emotion in animal models over the past two decades has implicated the amygdala in fear and other emotional processes. This work stimulated interest in pursuing the brain mechanisms of emotion in humans. Here, we review research on the role of the amygdala in emotional processes in both animal models and humans. The review is not exhaustive, but it highlights five major research topics that illustrate parallel roles for the amygdala in humans and other animals, including implicit emotional learning and memory, emotional modulation of memory, emotional influences on attention and perception, emotion and social behavior, and emotion inhibition and regulation.

ARTICLE UPDATE - Segregated neural representation of distinct emotion dimensions in the prefrontal cortex—an fMRI study

Simone Grimm, Conny F. Schmidt, Felix Bermpohl, Alexander Heinzel, Yuliya Dahlem, Michael Wyss, Daniel Hell, Peter Boesiger, Heinz Boeker and Georg Northoff

NeuroImage, in press

Emotions are frequently characterized by distinct dimensions such as valence, intensity, and recognition. However, the exact neural representation of these dimensions in different prefrontal cortical regions remains unclear. One of the problems in revealing prefrontal cortical representation is that the very same regions are also involved in cognitive functions associated with emotion processing. We therefore conducted an fMRI study involving the viewing of emotional pictures (using the International Affective Picture System; IAPS) and controlled for associated cognitive processing like judgment and preceding attention. Functional activation was correlated with subjective post-scanning ratings of valence, intensity, and recognition. Valence significantly correlated with the functional response in ventromedial prefrontal cortex (VMPFC) and dorsolateral prefrontal cortex (DLPFC), intensity with activation in ventrolateral prefrontal cortex (VLPFC) and dorsomedial prefrontal cortex (DMPFC), and recognition with the functional response in perigenual anterior cingulate cortex (PACC). In conclusion, our results indicate segregated neural representation of the different emotion dimensions in different prefrontal cortical regions.

ARTICLE UPDATE - Neural mechanism for judging the appropriateness of facial affect

Ji-Woong Kim, Jae-Jin Kim, Bum Seok Jeong, Seon Wan Ki, Dong-Mi Im, Soo Jung Lee and Hong Shick Lee

Cognitive Brain Research, in press

Questions regarding the appropriateness of facial expressions in particular situations arise ubiquitously in everyday social interactions. To determine the appropriateness of facial affect, first of all, we should represent our own or the other's emotional state as induced by the social situation. Then, based on these representations, we should infer the possible affective response of the other person. In this study, we identified the brain mechanism mediating special types of social evaluative judgments of facial affect in which the internal reference is related to theory of mind (ToM) processing. Many previous ToM studies have used non-emotional stimuli, but, because so much valuable social information is conveyed through nonverbal emotional channels, this investigation used emotionally salient visual materials to tap ToM. Fourteen right-handed healthy subjects volunteered for our study. We used functional magnetic resonance imaging to examine brain activation during the judgmental task for the appropriateness of facial affects as opposed to gender matching tasks. We identified activation of a brain network, which includes both medial frontal cortex, left temporal pole, left inferior frontal gyrus, and left thalamus during the judgmental task for appropriateness of facial affect compared to the gender matching task. The results of this study suggest that the brain system involved in ToM plays a key role in judging the appropriateness of facial affect in an emotionally laden situation. In addition, our result supports that common neural substrates are involved in performing diverse kinds of ToM tasks irrespective of perceptual modalities and the emotional salience of test materials.

ARTICLE UPDATE - How positive affect modulates cognitive control: The costs and benefits of reduced maintenance capability.

Dreisbach G

Brain and Cognition, in press

Adaptive action in a constantly changing environment requires the ability to maintain intentions and goals over time and to flexibly switch between these goals in response to significant changes. argued that positive affect modulates these antagonistic control demands in favor of a more flexible but also more distractible behavior. In the present paper, the author will present further evidence for the affective modulation of cognitive control: mild positive affect reduced maintenance capability in a simple cuing paradigm (the AX Continuous Performance Task) as compared to negative and neutral affect. This reduced maintenance capability results in costs when a to be maintained goal has to be executed and conversely results in benefits when a to be maintained goal unexpectedly changes. The data will be discussed with respect to existing theories on positive affect, cognitive control, and dopamine.

ARTICLE UPDATE - Bridging emotion theory and neurobiology through dynamic systems modeling

Lewis MD.

Behavioral and Brain Sciences, 28, 169-245.

Efforts to bridge emotion theory with neurobiology can be facilitated by dynamic systems (DS) modeling. DS principles stipulate higher-order wholes emerging from lower-order constituents through bidirectional causal processes--offering a common language for psychological and neurobiological models. After identifying some limitations of mainstream emotion theory, I apply DS principles to emotion-cognition relations. I then present a psychological model based on this reconceptualization, identifying trigger, self-amplification, and self-stabilization phases of emotion-appraisal states, leading to consolidating traits. The article goes on to describe neural structures and functions involved in appraisal and emotion, as well as DS mechanisms of integration by which they interact. These mechanisms include nested feedback interactions, global effects of neuromodulation, vertical integration, action-monitoring, and synaptic plasticity, and they are modeled in terms of both functional integration and temporal synchronization. I end by elaborating the psychological model of emotion-appraisal states with reference to neural processes.

ARTICLE UPDATE - Modulation of retrieval processing reflects accuracy of emotional source memory.

Smith AP, Henson RN, Rugg MD, Dolan RJ.

Learning & Memory, 12, 427-429.

There is considerable evidence that encoding and consolidation of memory are modulated by emotion, but the retrieval of emotional memories is not well characterized. Here we manipulated the emotional context with which affectively neutral stimuli were associated during encoding, allowing us to examine neural activity associated with retrieval of emotional memories without confounding the emotional attributes of cue items and the retrieved context. Using a source memory procedure we were also able to compare how retrieval processing was modulated when the emotional encoding context was recollected or not. An interaction between emotional encoding context and accuracy of source memory revealed that successful retrieval of emotional context was associated with activity in left amygdala, and a left frontotemporal network including anterior insula, prefrontal cortex and cingulate. In contrast, when contextual retrieval was unsuccessful, items encoded in emotional contexts elicited enhanced activity in right amygdala and a right-lateralized network that included extrastriate visual areas. These findings indicate distinct effects of emotion on successful and unsuccessful retrieval of source information, including lateralization of amygdala responses.

ARTICLE UPDATE - Analysis of Single-Unit Responses to Emotional Scenes in Human Ventromedial Prefrontal Cortex

Hiroto Kawasaki, Ralph Adolphs, Hiroyuki Oya, Christopher Kovach, Hanna Damasio, Olaf Kaufman and Matthew Howard, III

Journal of Cognitive Neuroscience, 17, 1509-1518.

Lesion and functional imaging studies in humans have shown that the ventral and medial prefrontal cortex is critically involved in the processing of emotional stimuli, but both of these methods have limited spatiotemporal resolution. Conversely, neurophysiological studies of emotion in nonhuman primates typically rely on stimuli that do not require elaborate cognitive processing. To begin bridging this gap, we recorded from a total of 267 neurons in the left and right orbital and anterior cingulate cortices of four patients who had chronically implanted depth electrodes for monitoring epilepsy. Peristimulus activity was recorded to standardized, complex visual scenes depicting neutral, pleasant, or aversive content. Recording locations were verified with postoperative magnetic resonance imaging. Using a conservative, multistep statistical evaluation, we found significant responses in 56 neurons; 16 of these were selective for only one emotion class, most often aversive. The findings suggest sparse and widely distributed processing of emotional value in the prefrontal cortex, with a predominance of responses to aversive stimuli.

ARTICLE UPDATE - Attentional Bias to Pictures of Fear-Relevant Animals in a Dot Probe Task

Lipp, O. V., & Derakshan, N.

Emotion, 5, 365-369.

Attentional bias to fear-relevant animals was assessed in 69 participants not preselected on self-reported anxiety with the use of a dot probe task showing pictures of snakes, spiders, mushrooms, and flowers. Probes that replaced the fear-relevant stimuli (snakes and spiders) were found faster than probes that replaced the non-fear-relevant stimuli, indicating an attentional bias in the entire sample. The bias was not correlated with self-reported state or trait anxiety or with general fearfulness. Participants reporting higher levels of spider fear showed an enhanced bias to spiders, but the bias remained significant in low scorers. The bias to snake pictures was not related to snake fear and was significant in high and low scorers. These results indicate preferential processing of fear-relevant stimuli in an unselected sample.

ARTICLE UPDATE - The Role of Fear-Relevant Stimuli in Visual Search: A Comparison of Phylogenetic and Ontogenetic Stimuli

Brosch, T., & Sharma, D.

Emotion, 5, 360-364.

It has been argued that phylogenetic fear-relevant stimuli elicit preattentive capture of attention. To distinguish between fear relevance and time of appearance in evolutionary history, the authors compare phylogenetic and ontogenetic fear-relevant and fear-irrelevant stimuli in a visual search task. The authors found no evidence for a special role of phylogenetic fear-relevant stimuli; it seems that fear relevance in general is more important than is the evolutionary age. The pattern of results indicates that attention toward threatening stimuli is mainly affected by a late component that prolongs the disengagement of attention.

ARTICLE UPDATE - Visual Search With Biological Threat Stimuli: Accuracy, Reaction Times, and Heart Rate Changes

Anders Flykt

Emotion, 5, 349-353.

Twenty-four participants were given a visual search task of deciding whether all the pictures in 3 × 3 search arrays contained a target picture from a deviant category, and heart rate was measured. The categories were snakes, spiders, flowers, and mushrooms. Shorter reaction times (RTs) were observed for fear-relevant (snake and spider) targets rather than for fear-irrelevant/neutral (flower and mushroom) targets. This difference was most pronounced for the participants presented with a gray-scale version of the search arrays. The 1st interbeat interval (IBI), after the search array onset, showed an effect of the target, whereas the 2nd IBI also showed an effect of the distractors. The results suggest that controlled processing of the task operates together with automatic processing.

ARTICLE UPDATE - Integrated neural representations of odor intensity and affective valence in human amygdala.

Winston JS, Gottfried JA, Kilner JM, Dolan RJ.

Journal of Neuroscience, 25, 8903-8907.

Arousal and valence are proposed to represent fundamental dimensions of emotion. The neural substrates for processing these aspects of stimuli are studied widely, with recent studies of chemosensory processing suggesting the amygdala processes intensity (a surrogate for arousal) rather than valence. However, these investigations have assumed that a valence effect in the amygdala is linear such that testing valence extremes is sufficient to infer responses across valence space. In this study, we tested an alternative hypothesis, namely that valence responses in the amygdala are nonlinear. Using event-related functional magnetic resonance imaging, we measured amygdala responses to high- and low-concentration variants of pleasant, neutral, and unpleasant odors. Our results demonstrate that the amygdala exhibits an intensity-by-valence interaction in olfactory processing. In other words, the effect of intensity on amygdala activity is not the same at all levels of valence. Specifically, the amygdala responds differentially to high (vs low)-intensity odor for pleasant and unpleasant smells but not for neutral smells. This implies that the amygdala codes neither intensity nor valence per se, but a combination that we suggest reflects the overall emotional value of a stimulus.

ARTICLE UPDATE - Emotion and Cognition: Insights from Studies of the Human Amygdala.

Phelps EA.

Annual Review of Psychology, 57, 27-53

Traditional approaches to the study of cognition emphasize an information- processing view that has generally excluded emotion. In contrast, the recent emergence of cognitive neuroscience as an inspiration for understanding human cognition has highlighted its interaction with emotion. This review explores insights into the relations between emotion and cognition that have resulted from studies of the human amygdala. Five topics are explored: emotional learning, emotion and memory, emotion's influence on attention and perception, processing emotion in social stimuli, and changing emotional responses. Investigations into the neural systems underlying human behavior demonstrate that the mechanisms of emotion and cognition are intertwined from early perception to reasoning. These findings suggest that the classic division between the study of emotion and cognition may be unrealistic and that an understanding of human cognition requires the consideration of emotion.

ARTICLE UPDATE - The somatic marker hypothesis: A critical evaluation

Dunn BD, Dalgleish T, Lawrence AD.

Neuroscience Biobehavior Review, in press

The somatic marker hypothesis (SMH; [Damasio, A. R., Tranel, D., Damasio, H., 1991. Somatic markers and the guidance of behaviour: theory and preliminary testing. In Levin, H.S., Eisenberg, H.M., Benton, A.L. (Eds.), Frontal Lobe Function and Dysfunction. Oxford University Press, New York, pp. 217-229]) proposes that emotion-based biasing signals arising from the body are integrated in higher brain regions, in particular the ventromedial prefrontal cortex (VMPFC), to regulate decision-making in situations of complexity. Evidence for the SMH is largely based on the performance on the Iowa Gambling Task (IGT; [Bechara, A., Tranel, D., Damasio, H., Damasio, A.R., 1996. Failure to respond autonomically to anticipated future outcomes following damage to prefrontal cortex. Cerebral Cortex 6 (2), 215-225]), linking anticipatory skin conductance responses (SCRs) to successful performance on a decision-making paradigm in healthy participants. These 'marker' signals were absent in patients with VMPFC lesions and were associated with poorer IGT performance. The current article reviews the IGT findings, arguing that their interpretation is undermined by the cognitive penetrability of the reward/punishment schedule, ambiguity surrounding interpretation of the psychophysiological data, and a shortage of causal evidence linking peripheral feedback to IGT performance. Further, there are other well-specified and parsimonious explanations that can equally well model the IGT data. Next, lesion, neuroimaging, and psychopharmacology data evaluating the proposed neural substrate underpinning the SMH are reviewed. Finally, conceptual reservations about the novelty, parsimony and specification of the SMH are raised. It is concluded that while presenting an elegant theory of how emotion influences decision-making, the SMH requires additional empirical support to remain tenable.

ARTICLE UPDATE - With Sadness Comes Accuracy; With Happiness, False Memory: Mood and the False Memory Effect

Storbeck, Justin; Clore, Gerald L.

Psychological Science, 16, 785-791.

The Deese-Roediger-McDermott paradigm lures people to produce false memories. Two experiments examined whether induced positive or negative moods would influence this false memory effect. The affect-as-information hypothesis predicts that, on the one hand, positive affective cues experienced as task-relevant feedback encourage relational processing during encoding, which should enhance false memory effects. On the other hand, negative affective cues are hypothesized to encourage item-specific processing at encoding, which should discourage such effects. The results of Experiment 1 are consistent with these predictions: Individuals in negative moods were significantly less likely to show false memory effects than those in positive moods or those whose mood was not manipulated. Experiment 2 introduced inclusion instructions to investigate whether moods had their effects at encoding or retrieval. The results replicated the false memory finding of Experiment 1 and provide evidence that moods influence the accessibility of lures at encoding, rather than influencing monitoring at retrieval of whether lures were actually presented.

ARTICLE UPDATE - Attentional Inhibition Has Social-Emotional Consequences for Unfamiliar Faces

Fenske, Mark J.; Raymond, Jane E.; Kessler, Klaus; Westoby, Nikki; Tipper, Steven P.

Psychological Science, 16, 753-758

Visual attention studies often rely on response time measures to show the impact of attentional facilitation and inhibition. Here we extend the investigation of the effects of attention on behavior and show that prior attentional states associated with unfamiliar faces can influence subsequent social-emotional judgments about those faces. Participants were shown pairs of face images and were asked to withhold a response if a transparent stop-signal cue appeared over one of the faces. This served to associate the cued face with an inhibitory state. Later, when asked to make social-emotional choices about these face pairs, participants chose uncued faces more often than cued faces as “more trustworthy” and chose cued faces more often than uncued faces as “less trustworthy.” For perceptual choices, there was no effect of how the question was framed (which face is “on a lighter background” vs. “on a darker background”). These results suggest that attentional inhibition can be associated with socially relevant stimuli, such as faces, and can have specific, deleterious effects on social-emotional judgments.

ARTICLE UPDATE - The role of spatial frequency information for ERP components sensitive to faces and emotional facial expression

Amanda Holmes, Joel S. Winston and Martin Eimer

Cognitive Brain Research, 25, 508-520

To investigate the impact of spatial frequency on emotional facial expression analysis, ERPs were recorded in response to low spatial frequency (LSF), high spatial frequency (HSF), and unfiltered broad spatial frequency (BSF) faces with fearful or neutral expressions, houses, and chairs. In line with previous findings, BSF fearful facial expressions elicited a greater frontal positivity than BSF neutral facial expressions, starting at about 150 ms after stimulus onset. In contrast, this emotional expression effect was absent for HSF and LSF faces. Given that some brain regions involved in emotion processing, such as amygdala and connected structures, are selectively tuned to LSF visual inputs, these data suggest that ERP effects of emotional facial expression do not directly reflect activity in these regions. It is argued that higher order neocortical brain systems are involved in the generation of emotion-specific waveform modulations. The face-sensitive N170 component was neither affected by emotional facial expression nor by spatial frequency information.

ARTICLE UPDATE - An emotional mediation theory of differential age effects in episodic and semantic memories

Allen PA, Kaut KP, Lord RG, Hall RJ, Grabbe JW, Bowie T.

Experimental Aging Research,31,355-91

Although there is a large decrement in central episodic memory processes as adults age, there is no appreciable decrement in central semantic memory processes (Allen et al., Journal of Gerontology: Psychological Sciences, 57B, P173-P186, 2002; Allen et al., Experimental Aging Research, 28, 111-142, 2002; Mitchell, Journal of Experimental Psychology: Learning, Memory, and Cognition, 15, 31-49, 1989). The authors develop a theory of episodic memory's connections to cognitive, emotional, and motivational systems to explain these differential age effects. The theory is discussed within the context of the cognitive neuroscience research regarding limbic system connectivity in conjunction with Damasio's notion of somatic markers (Descartes' error: Emotion, reason, and the human brain, New York: Grosset/Putnam, 1994). The central hypothesis is that elements of limbic system circuitry, including portions of the medial temporal lobes and frontal cortex, are associated with both working and long-term episodic memory performance, and by extension, with the capacity to engage in emotion-guided, self-regulatory processes that depend heavily on episodic memory. In contrast, the semantic memory system may have less shared interface with episodic and affective networks (i.e., the limbic-related system), and therefore remain independent of neurocognitive changes impacting emotional states and episodic-type memory processes. Accordingly, this framework may account for the pattern of age-related declines in episodic relative to semantic memory, particularly if older adults experience less emotional activation, and therefore fewer somatic markers, than younger adults. An initial empirical examination of this emotional mediation theory is presented, using preexisting data that include indicators of age, chronic tendency to focus on negative emotional stimuli (neuroticism), and working memory performance.

ARTICLE UPDATE - Emotional enhancement of perceptual priming is preserved in aging and early-stage Alzheimer's disease

Kevin S. LaBar, Dana C. Torpey, Craig A. Cook, Stephanie R. Johnson, Lauren H. Warren, James R. Burke and Kathleen A. Welsh-Bohmer

Neuropsychologica, 43, 1824-1837.

Perceptual priming for emotionally-negative and neutral scenes was tested in early-stage Alzheimer's disease (AD) patients and healthy younger, middle-aged and older adults. In the study phase, participants rated the scenes for their arousal properties. In the test phase, studied and novel scenes were initially presented subliminally, and the exposure duration was gradually increased until a valence categorization was made. The difference in exposure duration required to categorize novel versus studied items was the dependent measure of priming. Aversive content increased the magnitude of priming, an effect that was preserved in healthy aging and AD. Results from an immediate recognition memory test showed that the priming effects could not be attributable to enhanced explicit memory for the aversive scenes. These findings implicate a dissociation between the modulatory effect of emotion across implicit and explicit forms of memory in aging and early-stage AD.

ARTICLE UPDATE - Empathy examined through the neural mechanisms involved in imagining how I feel versus how you feel pain

Philip L. Jackson, Eric Brunet, Andrew N. Meltzoff and Jean Decety

Neuropsychologia, in press

Perspective-taking is a stepping stone to human empathy. When empathizing with another individual, one can imagine how the other perceives the situation and feels as a result. To what extent does imagining the other differs from imagining oneself in similar painful situations? In this functional magnetic resonance imaging experiment, participants were shown pictures of people with their hands or feet in painful or non-painful situations and instructed to imagine and rate the level of pain perceived from different perspectives. Both the Self's and the Other's perspectives were associated with activation in the neural network involved in pain processing, including the parietal operculum, anterior cingulate cortex (ACC; BA32) and anterior insula. However, the Self-perspective yielded higher pain ratings and involved the pain matrix more extensively in the secondary somatosensory cortex, the ACC (BA 24a′/24b′), and the insula proper. Adopting the perspective of the Other was associated with specific increase in the posterior cingulate/precuneus and the right temporo-parietal junction. These results show the similarities between Self- and Other-pain representation, but most interestingly they also highlight some distinctiveness between these two representations, which is a crucial aspect of human empathy. It may be what allows us to distinguish empathic responses to others versus our own personal distress. These findings are consistent with the view that empathy does not involve a complete Self–Other merging.

ARTICLE UPDATE - Functional dissociation of amygdala-modulated arousal and cognitive appraisal, in Turner syndrome

D. H. Skuse, J. S. Morris and R. J. Dolan

Brain, 128, 2084-2096

The amygdala is preferentially activated by facial expressions of fear. Right and left amygdala are hypothesized to play distinct, but complementary, roles that influence somatic and cognitive responses to facial expressions. Right amygdala activation is linked to autonomic arousal, and thus indirectly influences left hemisphere cognitive processing centres. Left amygdala activation is more closely associated with cognitive processing and differentiation of facial emotions. A double-dissociation between the functions of left and right amygdala is implied by lesion studies but supportive evidence is inconsistent, partly because patients with structural anteromedial temporal anomalies have experienced variable surgical procedures. A functional dissociation can be demonstrated between arousal and the cognitive appraisal of fearful faces in the condition of X-monosomy or Turner syndrome. Previous research found Turner syndrome women of normal verbal intelligence are seriously impaired in their ability cognitively to differentiate fearful from other facial expressions but they acquire fear conditioning normally, with enhanced autonomic responses. These findings supported the dissociation hypothesis, which was formally tested in a study of 12 X-monosomic and 12 control females who participated in functional magnetic resonance imaging during which simultaneous skin conductance recordings were acquired. Faces depicting fear or neutral emotions were presented to both case and control subjects in random order. Arousal to (fearful–neutral) faces was associated with transiently increased skin conductance responses and bilateral amygdala activation in both groups, but X-monosomic females had proportionately greater—and more persistent—right amygdala activation than controls. In both groups, cognitive accuracy correlated positively with differential activity of left fusiform gyrus. There was a significant correlation between the left fusiform and left medial amygdala activation only in normal females, and only in them did differential SCRs (to fearful–neutral faces) correlate positively with left fusiform responses. Arousal and cognitive appraisal functions of the amygdala can thus be functionally dissociated. X-monosomy selectively impairs explicit recognition of fearful faces in the presence of normal or enhanced autonomic reactivity, and is associated with a functional dissociation of activity in left amygdala and left fusiform gyrus. These findings imply X-linked genes are essential for binding somatic responses to the cognitive appraisal of emotional stimuli.

ARTICLE UPDATE - Neural evidence of effects of emotional valence on word recognition

Midori Inaba, Michio Nomura and Hideki Ohira

International Journal of Psychophysiology, 57, 165-173

There are no clear reports of electrophysiological evidence of the facilitating effect of negative valence on word recognition. However, behavioral psychological studies have suggested that negative words can be recognized more accurately than positive and neutral words. This study aimed to examine whether, and if so how, the valence of words could influence accuracy and event-related potentials (ERPs) in a recognition task. ERPs were recorded from 20 healthy subjects during performance of a word recognition task. We found a behavioral advantage in discriminability between old and new items for negative words. As for ERPs, the positive-going shift was evident for correct responses to targets in late latency at midline and left centro-parietal sites. Additionally, the magnitude of this component was greatest for negative targets, next for positive targets, and least for neutral targets. The findings offer support for the idea that negative content greatly accelerates recognition memory compared to positive and neutral words.

ARTICLE UPDATE - Memory for specific visual details can be enhanced by negative arousing content

Elizabeth A. Kensinger, Rachel J. Garoff-Eaton and Daniel L. Schacter

Journal of Memory and Language, 54(1), 99-112

Individuals often claim that they vividly remember information with negative emotional content. At least two types of information could lead to this sense of enhanced vividness: Information about the emotional item itself (e.g., the exact visual details of a snake) and information about the context in which the emotional item was encountered (e.g., the fact that the snake was sitting on a branch in a forest). The present study focused on the former, investigating how exposure duration at study and emotional content of an object affected the likelihood of remembering an item’s specific visual details. Participants studied neutral objects (e.g., a barometer) and negative arousing objects (e.g., a grenade) and were later shown either the identical object or a different photo of the same type of object (e.g., another barometer). Across two experiments, emotional content enhanced the likelihood that specific visual details were remembered: Individuals were more likely to correctly indicate that an item was identical to the object studied earlier if it was an emotional object than if it was a neutral object. This memory benefit for the emotional items was most robust when items were shown for longer exposure durations (500 or 1000 ms) rather than only briefly (for 250 ms). Thus, with sufficient processing time, negative arousing content appears to enhance the likelihood that visual details are remembered about an object.

ARTICLE UPDATE - Differential Effects of Lesions of the Amygdala and Prefrontal Cortex on Recognizing Facial Expressions of Complex Emotions

P. Shaw, J. Bramham, E. J. Lawrence, R. Morris, S. Baron-Cohen and A. S. David

Journal of Cognitive Neuroscience, 17, 1410-1419.

Humans can detect facial expressions of both simple, basic emotions and expressions reflecting more complex states of mind. The latter includes emotional expressions that regulate social interactions ("social expressions" such as looking hostile or friendly) and expressions that reflect the inner thought state of others ("cognitive expressions" such as looking pensive). To explore the neural substrate of this skill, we examined performance on a test of detection of such complex expressions in patients with lesions of the temporal lobe (n = 54) or frontal lobe (n = 31). Of the temporal group, 18 had unilateral focal lesions of the amygdala and of the frontal group, 14 patients had unilateral lesions of the ventromedial prefrontal cortex—two regions held to be pivotal in mediating social cognitive skills. Damage to either the left or right amygdala was associated with impairment in the recognition of both social and cognitive expressions, despite an intact ability to extract information relating to invariant physical attributes. Lesions to all of the right prefrontal cortex—not just the ventromedial portions—led to a specific deficit in recognizing complex social expressions with a negative valence. The deficit in the group with right prefrontal cortical damage may contribute to the disturbances in social behavior associated with such lesions. The results also suggest that the amygdala has a role in processing a wide range of emotional expressions.

ARTICLE UPDATE - Unattended Facial Expressions Asymmetrically Bias the Concurrent Processing of Nonemotional Information

Jeffrey S. Maxwell, Alexander J. Shackman and Richard J. Davidson

Journal of Cognitive Neuroscience, 17, 1386-1395.

Planned and reflexive behaviors often occur in the presence of emotional stimuli and within the context of an individual's acute emotional state. Therefore, determining the manner in which emotion and attention interact is an important step toward understanding how we function in the real world. Participants in the current investigation viewed centrally displayed, task-irrelevant, face distractors (angry, neutral, happy) while performing a lateralized go/no-go continuous performance task. Lateralized go targets and no-go lures that did not spatially overlap with the faces were employed to differentially probe processing in the left (LH) and right (RH) cerebral hemispheres. There was a significant interaction between expression and hemisphere, with an overall pattern such that angry distractors were associated with relatively more RH inhibitory errors than neutral or happy distractors and happy distractors with relatively more LH inhibitory errors than angry or neutral distractors. Simple effects analyses confirmed that angry faces differentially interfered with RH relative to LH inhibition and with inhibition in the RH relative to happy faces. A significant three-way interaction further revealed that state anxiety moderated relations between emotional expression and hemisphere. Under conditions of low cognitive load, more intense anxiety was associated with relatively greater RH than LH impairment in the presence of both happy and threatening distractors. By contrast, under high load, only angry distractors produced greater RH than LH interference as a function of anxiety.

ARTICLE UPDATE - The neurobiology of positive emotions

Burgdorf J, Panksepp J.

Neuroscience & Biobehavioral Reviews, in press

Compared to the study of negative emotions such as fear, the neurobiology of positive emotional processes and the associated positive affect (PA) states has only recently received scientific attention. Biological theories conceptualize PA as being related to (i) signals indicating that bodies are returning to equilibrium among those studying homeostasis, (ii) utility estimation among those favoring neuroeconomic views, and (iii) approach and other instinctual behaviors among those cultivating neuroethological perspectives. Indeed, there are probably several distinct forms of positive affect, but all are closely related to ancient sub-neocortical limbic brain regions we share with other mammals. There is now a convergence of evidence to suggest that various regions of the limbic system, including especially ventral striatal dopamine systems are implemented in an anticipatory (appetitive) positive affective state. Dopamine independent mechanisms utilizing opiate and GABA receptors in the ventral striatum, amygdala and orbital frontal cortex are important in elaborating consummatory PA (i.e. sensory pleasure) states, and various neuropeptides mediate homeostatic satisfactions.

ARTICLE UPDATE - On the role of attention for the processing of emotions in speech: Sex differences revisited.

Schirmer A, Kotz SA, Friederici AD.

Cognitive Brain Research, 24, 442-52.

In a previous cross-modal priming study [A. Schirmer, A.S. Kotz, A.D. Friederici, Sex differentiates the role of emotional prosody during word processing, Cogn. Brain Res. 14 (2002) 228-233.], we found that women integrated emotional prosody and word valence earlier than men. Both sexes showed a smaller N400 in the event-related potential to emotional words when these words were preceded by a sentence with congruous compared to incongruous emotional prosody. However, women showed this effect with a 200-ms interval between prime sentence and target word whereas men showed the effect with a 750-ms interval. The present study was designed to determine whether these sex differences prevail when attention is directed towards the emotional content of prosody and word meaning. To this end, we presented the same prime sentences and target words as in our previous study. Sentences were spoken with happy or sad prosody and followed by a congruous or incongruous emotional word or pseudoword. The interval between sentence offset and target onset was 200 ms. In addition to performing a lexical decision, participants were asked to decide whether or not a word matched the emotional prosody of the preceding sentence. The combined lexical and congruence judgment failed to reveal differences in emotional-prosodic priming between men and women. Both sexes showed smaller N400 amplitudes to emotionally congruent compared to incongruent words. This suggests that the presence of sex differences in emotional-prosodic priming depends on whether or not participants are instructed to take emotional prosody into account.

ARTICLE UPDATE - Incidental effects of emotional valence in single word processing: An fMRI study

Lars Kuchinke, Arthur M. Jacobs, Claudia Grubich, Melissa L.-H. Võ, Markus Conrada and Manfred Herrmann

NeuroImage, 28, 1022-1032

The present study aimed at identifying the neural responses associated with the incidental processing of the emotional valence of single words using event-related functional magnetic resonance imaging (fMRI). Twenty right-handed participants performed a visual lexical decision task, discriminating between nouns and orthographically and phonologically legal nonwords. Positive, neutral and negative word categories were matched for frequency, number and frequency of orthographic neighbors, number of letters and imageability. Response times and accuracy data differed significantly between positive and neutral, and positive and negative words respectively, thus, replicating the findings of a pilot study. Words showed distributed, mainly left hemisphere activations, indicating involvement of a neural network responsible for semantic word knowledge. The neuroimaging data further revealed areas in left orbitofrontal gyrus and bilateral inferior frontal gyrus with greater activation to emotional than to neutral words. These brain regions are known to be involved in processing semantic and emotional information. Furthermore, distinct activations associated with positive words were observed in bilateral middle temporal and superior frontal gyrus, known to support semantic retrieval, and a distributed network, namely anterior and posterior cingulate gyrus, lingual gyrus and hippocampus when comparing positive and negative words. The latter areas were previously associated with explicit and not incidental processing of the emotional meaning of words and emotional memory retrieval. Thus, the results are discussed in relation to models of processing semantic and episodic emotional information.

ARTICLE UPDATE - Functional consequences of perceiving facial expressions of emotion without awareness.

Eastwood JD, Smilek D.

Consciousness & Cognition, 14, 565-84

A substantial body of research has established that even when we are not consciously aware of the faces of others we are nevertheless sensitive to, and impacted by their facial expression. In this paper, we consider this body of research from a new perspective by examining the functions of unconscious perception revealed by these studies. A consideration of the literature from this perspective highlights that existing research methods are limited when it comes to revealing possible functions of unconscious perception. The critical shortcoming is that in all of the methods, the perceived facial expression remains outside of awareness. This is a problem because there are good reasons to believe that one important function of unconsciously perceived negative faces is to attract attention so that they are consciously perceived; such conscious perception, however, is never allowed with existing methodologies. We discuss recent studies of emotional face perception under conditions of visual search that address this issue directly. Further, we suggest that methodologies that do not examine cognitive processes as they occur in more natural settings may result in fundamental misunderstandings of human cognition.

ARTICLE UPDATE - How do we modulate our emotions? Parametric fMRI reveals cortical midline structures as regions specifically involved in the processi

Alexander Heinzel, Felix Bermpohl, Robert Niese, Andrea Pfennig, Alvaro Pascual-Leone, Gottfried Schlaug and Georg Northoff

Cognitive Brain Research, 25,348-358.

One of the major problems in affective neuroscience of healthy subjects as well as of patients with emotional dysfunctions is to disentangle emotional core functions and non-emotional processes. Emotional valence is considered an emotional key process. The present study employed a parametric functional magnetic resonance imaging (fMRI) study to address this question. Thirteen healthy volunteers were scanned during emotional stimulus processing (International Affective Picture System). The presented pictures covered the entire range of emotional valences. The fMRI data were consecutively subjected to a preliminary categorical (valence-independent) and a detailed parametric analysis, the latter using individual valence ratings as regressor. The parametric analysis revealed a linear valence-dependent modulation of the BOLD signal in the orbito- and dorsomedial prefrontal cortex (OMPFC, DMPFC), medial parietal cortex (MPC), and insula. In addition, we observed that emotional valence exerts its effects predominantly via modulation of signal decreases. We conclude that the psychological concept of emotional valence may be related to neural processing in cortical midline regions.

ARTICLE UPDATE - Signals for threat modulate attentional capture and holding: Fear‐conditioning and extinction during the exogenous cueing task

Ernst H. W. Koster , Geert Crombez , Stefaan Van Damme , Bruno Verschuere , Jan De Houwer

Cognition & Emotion, 18, 771-780.

This experiment investigated whether acquired signals for threat capture and hold visual attention. Cues that were presented in an exogenous attentional cueing task were emotionally modulated using a fear‐conditioning paradigm. During acquisition, undergraduate students (55 women, 11 men) learned that one cue (CS+) of the attentional task was a signal for an aversive white noise burst (UCS, 100 dBA) and that another cue (CS−) signalled its nonoccurrence. In a subsequent extinction phase, no UCSs were presented anymore during the cueing task. Results indicated that during acquisition, the CS+ cues strongly captured and held visual attention in comparison with the CS− cues, and that these effects diminished during extinction.

ARTICLE UPDATE - Affective priming as an indirect measure of food preferences acquired through odor conditioning.

Hermans D, Baeyens F, Lamote S, Spruyt A, Eelen P.

Experimental Psychology,52, 180-186.

The present study aimed at investigating affective priming for originally neutral food stimuli that recently acquired their affective meaning through odor conditioning. In a first phase, pictures of different brands of yogurts (CSs) were contingently presented with a positive or negative odor (US). In a subsequent phase, the yogurt CSs were used as primes in an affective priming procedure. Rating data showed that the acquisition procedure resulted in a reliable evaluative learning effect. This could be corroborated by the results of the priming task. Participants responded faster to positive target words and made fewer errors when they were preceded by a CS that had been associated with a positive odor, as compared to a CS that was associated with a negative odor. A reversed pattern was present for negative targets. Based on these findings, it is suggested that affective priming might be used as a demand-free measure of evaluative learning.

ARTICLE UPDATE - When less is more: the consequences of affective primacy for subliminal priming effects.

Stapel DA, Koomen W.

Personality & Socical Psychological Bulletin, 31, 1286-1295.

This research investigates the consequences of the notion that one can distinguish early-evaluative (when exposure is short) and late-descriptive reactions (when exposure is long) to subliminally primed trait concepts. In three studies, it was found that the evaluative effects instigated by short exposure to primed concepts were bigger than the evaluative + descriptive effects instigated by long exposure: Less is more. Only when exposure was short, target interpretations were accompanied by evaluative inferences (Studies 1 and 3). Similarly, only when exposure was short, descriptively inapplicable trait primes affected the interpretation of an ambiguous target (Studies 2 and 3).

ARTICLE UPDATE - Cognitive-emotional distinctiveness: Separating emotions from non-emotions in the representation of a stressful memory.

Boals A, Klein K.

Memory, 13,638-648.

Current theories on autobiographical memory and recent neurological evidence suggest that emotional and non-emotional features of a memory may be retrieved by separate systems. To test this notion, 207 participants who had experienced the break-up of a significant romantic relationship in the last 12 months completed a Multidimensional Scaling (MDS) procedure in relation to the previous relationship. The resulting MDS model revealed two dimensions: a valence and an emotional/non-emotional dimension. Further, participants who associated a high level of distress with their relationship break-up perceived less dissimilarity between emotional and non-emotional features than participants who associated a low level of distress with their relationship break-up. Theoretical and methodological implications for stress and memory are discussed.

ARTICLE UPDATE - Investigating emotion with music: An fMRI study.

Koelsch S, Fritz T, V Cramon DY, Muller K, Friederici AD.

Human Brain Mapping, in press

The present study used pleasant and unpleasant music to evoke emotion and functional magnetic resonance imaging (fMRI) to determine neural correlates of emotion processing. Unpleasant (permanently dissonant) music contrasted with pleasant (consonant) music showed activations of amygdala, hippocampus, parahippocampal gyrus, and temporal poles. These structures have previously been implicated in the emotional processing of stimuli with (negative) emotional valence; the present data show that a cerebral network comprising these structures can be activated during the perception of auditory (musical) information. Pleasant (contrasted to unpleasant) music showed activations of the inferior frontal gyrus (IFG, inferior Brodmann's area (BA) 44, BA 45, and BA 46), the anterior superior insula, the ventral striatum, Heschl's gyrus, and the Rolandic operculum. IFG activations appear to reflect processes of music-syntactic analysis and working memory operations. Activations of Rolandic opercular areas possibly reflect the activation of mirror-function mechanisms during the perception of the pleasant tunes. Rolandic operculum, anterior superior insula, and ventral striatum may form a motor-related circuitry that serves the formation of (premotor) representations for vocal sound production during the perception of pleasant auditory information. In all of the mentioned structures, except the hippocampus, activations increased over time during the presentation of the musical stimuli, indicating that the effects of emotion processing have temporal dynamics; the temporal dynamics of emotion have so far mainly been neglected in the functional imaging literature.

ARTICLE UPDATE - Unintentional processing of motivational valence

Agnes Moors, Jan De Houwer, Dirk Hermans, Paul Eelen

The Quarterly Journal of Experimental Psychology: Section A, 58, 1043-1063

Recent motivational affective priming studies (Moors & De Houwer, 2001; Moors, De Houwer, & Eelen, 2004) showed that primes that indicate success on a goal-inducing task facilitate positive target responses whereas primes that indicate failure on that task facilitate negative target responses. In the current studies, we examined whether these priming effects depend on consciously intentional processing of the motivational valence of the primes. In Experiment 1, the outcome of success or failure was presented not only immediately before the target (i.e., the prime) but also a second time after the target response. This should encourage participants to ignore the prime. In Experiment 2, participants were asked to respond to the targets within 600 ms after target onset. As a result, participants had little opportunity to process the motivational prime valence in a consciously intentional way. Nevertheless, strong affective priming effects were found in both studies. These results provide additional support for the claim that motivational valence can be processed without the conscious intention to do so.

ARTICLE UPDATE - Emotion and attention interactions in social cognition: Brain regions involved in processing anger prosody

David Sander, Didier Grandjean, Gilles Pourtois, Sophie Schwartz, Mohamed L. Seghier, Klaus R. Scherer and Patrik Vuilleumier

NeuroImage, 28,848-858

Multiple levels of processing are thought to be involved in the appraisal of emotionally relevant events, with some processes being engaged relatively independently of attention, whereas other processes may depend on attention and current task goals or context. We conducted an event-related fMRI experiment to examine how processing angry voice prosody, an affectively and socially salient signal, is modulated by voluntary attention. To manipulate attention orthogonally to emotional prosody, we used a dichotic listening paradigm in which meaningless utterances, pronounced with either angry or neutral prosody, were presented simultaneously to both ears on each trial. In two successive blocks, participants selectively attended to either the left or right ear and performed a gender-decision on the voice heard on the target side. Our results revealed a functional dissociation between different brain areas. Whereas the right amygdala and bilateral superior temporal sulcus responded to anger prosody irrespective of whether it was heard from a to-be-attended or to-be-ignored voice, the orbitofrontal cortex and the cuneus in medial occipital cortex showed greater activation to the same emotional stimuli when the angry voice was to-be-attended rather than to-be-ignored. Furthermore, regression analyses revealed a strong correlation between orbitofrontal regions and sensitivity on a behavioral inhibition scale measuring proneness to anxiety reactions. Our results underscore the importance of emotion and attention interactions in social cognition by demonstrating that multiple levels of processing are involved in the appraisal of emotionally relevant cues in voices, and by showing a modulation of some emotional responses by both the current task-demands and individual differences.

ARTICLE UPDATE - A critical period for the impact of amygdala damage on the emotional enhancement of memory?

P. Shaw, MD, B. Brierley, PhD and A. S. David, MD

Neurology, 65, 326-328

The amygdala is crucial in modulating enhanced memory for emotionally arousing material. The authors provide evidence that unilateral lesions of the human amygdala arising early in development, but not in adulthood, are associated with a loss of the expected superior retrieval of emotionally arousing over neutral material. This adds to evidence for an early critical period in the development of amygdala function.

ARTICLE UPDATE - Dissociation of emotional processes in response to visual and olfactory stimuli following frontotemporal damage

Soussignan R, Ehrle N, Henry A, Schaal B, Bakchine S.

Neurocase, 11, 114-28

Contemporary neuropsychological studies have stressed the widely distributed and multicomponential nature of human affective processes. Here, we examined facial electromyographic (EMG) (zygomaticus and corrugator muscle activity), autonomic (skin conductance and heart rate) and subjective measures of affective valence and arousal in patient TG, a 30 year-old man with left anterior mediotemporal and left orbitofrontal lesions resulting from a traumatic brain injury. Both TG and a normal control group were exposed to hedonically valenced visual and olfactory stimuli. In contrast with control subjects, facial EMG and electrodermal activity in TG did not differentiate among pleasant, unpleasant and neutral pictures. In addition, the controls reacted spontaneously with larger corrugator EMG activity and higher skin conductance to unpleasant odors. By contrast, the subjective feeling states (pleasure and arousal ratings) remained preserved in TG. The covariation between facial and self-report measures of negative valence was also a function of the nature of the olfactory task in the patient only. Taken together, the data suggest a functional dissociation between brain substrates supporting generation of emotion and those supporting representation of emotion.

ARTICLE UPDATE - Attention and emotion: Does rating emotion alter neural responses

C.A. Hutcherson, P.R. Goldin, K.N. Ochsner, J.D. Gabrieli,
L. Feldman Barrett, and J.J. Grossa

NeuroImage, in press

Functional neuroimaging of affective systems often includes subjective self-report of the affective response. Although self-report provides valuable information regarding participants’ affective responses, prior studies have raised the concern that the attentional demands of reporting on affective experience may obscure neural activations reflecting more natural affective responses. In the present study, we used potent emotion-eliciting amusing and sad films, employed a novel method of continuous self-reported rating of emotion experience, and compared the impact of rating with passive viewing of amusing and sad films. Subjective rating of ongoing emotional responses did not decrease either self-reported experience of emotion or neural activations relative to passive viewing in any brain regions. Rating, relative to passive viewing, produced increased activity in anterior cingulate, insula, and several other areas associated with introspection of emotion. These results support the use of continuous emotion measures and emotionally engaging films to study the dynamics of emotional responding and suggest that there may be some contexts in which the attention to emotion induced by reporting emotion experience does not disrupt emotional responding either behaviorally or neurally.

ARTICLE UPDATE - Attitudes to the right- and left: Frontal ERP asymmetries associated with stimulus valence and processing goals

William A. Cunningham, Stacey D. Espinet, Colin G. DeYoung and Philip David Zelazo

NeuroImage, 28, 827-834

We used dense-array event-related potentials (ERP) to examine the time course and neural bases of evaluative processing. Participants made good vs. bad (evaluative) and abstract vs. concrete (nonevaluative) judgments of socially relevant concepts (e.g., “murder,” “welfare”), and then rated all concepts for goodness and badness. Results revealed a late positive potential (LPP) beginning at about 475 ms post-stimulus and maximal over anterior sites. The LPP was lateralized (higher amplitude and shorter latency) on the right for concepts later rated bad, and on the left for concepts later rated good. Moreover, the degree of lateralization for the amplitude but not the latency was larger when participants were making evaluative judgments than when they were making nonevaluative judgments. These data are consistent with a model in which discrete regions of prefrontal cortex (PFC) are specialized for the evaluative processing of positive and negative stimuli.

ARTICLE UPDATE - From emotion perception to emotion experience: Emotions evoked by pictures and classical music.

Baumgartner T, Esslen M, Jancke L.

International Journal of Psychophysiology, in press

Most previous neurophysiological studies evoked emotions by presenting visual stimuli. Models of the emotion circuits in the brain have for the most part ignored emotions arising from musical stimuli. To our knowledge, this is the first emotion brain study which examined the influence of visual and musical stimuli on brain processing. Highly arousing pictures of the International Affective Picture System and classical musical excerpts were chosen to evoke the three basic emotions of happiness, sadness and fear. The emotional stimuli modalities were presented for 70 s either alone or combined (congruent) in a counterbalanced and random order. Electroencephalogram (EEG) Alpha-Power-Density, which is inversely related to neural electrical activity, in 30 scalp electrodes from 24 right-handed healthy female subjects, was recorded. In addition, heart rate (HR), skin conductance responses (SCR), respiration, temperature and psychometrical ratings were collected. Results showed that the experienced quality of the presented emotions was most accurate in the combined conditions, intermediate in the picture conditions and lowest in the sound conditions. Furthermore, both the psychometrical ratings and the physiological involvement measurements (SCR, HR, Respiration) were significantly increased in the combined and sound conditions compared to the picture conditions. Finally, repeated measures ANOVA revealed the largest Alpha-Power-Density for the sound conditions, intermediate for the picture conditions, and lowest for the combined conditions, indicating the strongest activation in the combined conditions in a distributed emotion and arousal network comprising frontal, temporal, parietal and occipital neural structures. Summing up, these findings demonstrate that music can markedly enhance the emotional experience evoked by affective pictures.

ARTICLE UPDATE - Repression of unconscious information by conscious processing: Evidence from affective blindsight induced by transcranial magnetic st

Jolij J, Lamme VA

PNAS, in press

Some patients with a lesion to the primary visual cortex (V1) show "blindsight": the remarkable ability to guess correctly about attributes of stimuli presented to the blind hemifield. Here, we show that blindsight can be induced in normal observers by using transcranial magnetic stimulation of the occipital cortex but exclusively for the affective content of unseen stimuli. Surprisingly, access to the affective content of stimuli disappears upon prolonged task training or when stimulus visibility increases, allegedly increasing the subjects' confidence in their overall performance. This finding suggests that availability of conscious information suppresses access to unconscious information, supporting the idea of consciousness as a repressant of unconscious tendencies.

ARTICLE UPDATE - Fate of unattended fearful faces in the amygdala is determined by both attentional resources and cognitive modulation

Luiz Pessoa, Srikanth Padmala and Thomas Morland

NeuroImage, in press

The evidence for amygdala processing of emotional items outside the focus of attention is mixed. We hypothesized that differences in attentional demands may, at least in part, explain prior discrepancies. In the present study, attention was manipulated by parametrically varying the difficulty of a central task, allowing us to compare responses evoked by unattended emotion-laden faces while the attentional load of a central task was varied. Reduced responses to unattended emotional stimuli may also reflect an active suppression of amygdala responses during difficult non-emotional tasks (cognitive modulation). To explicitly assess cognitive modulation, an experimental condition was used in which subjects performed the central task without the presence of irrelevant emotional stimuli. Our findings revealed that amygdala responses were modulated by the focus of attention. Stronger responses were evoked during a sex task (when faces were attended) relative to a bar-orientation task (when faces were unattended). Critically, a valence effect was observed in the right amygdala during low attentional demand conditions, but not during medium or high demand conditions. Moreover, performing a difficult non-emotional task alone was associated with signal decreases in a network of brain regions, including the amygdala. Such robust decreases demonstrate that cognitive modulation comprises a powerful factor in determining amygdala responses. Collectively, our findings reveal that both attentional resources and cognitive modulation govern the fate of unattended fearful faces in the amygdala.

ARTICLE UPDATE - Placebo in Emotional Processing— Induced Expectations of Anxiety Relief Activate a Generalized Modulatory Network

Predrag Petrovic, Thomas Dietrich, Peter Fransson, Jesper Andersson, Katrina Carlsson and Martin Ingvar

Neuron, 46, 957-969

Placebo analgesia and reward processing share several features. For instance, expectations have a strong influence on the subsequent emotional experience of both. Recent imaging data indicate similarities in the underlying neuronal network. We hypothesized that placebo analgesia is a special case of reward processing and that placebo treatment could modulate emotional perception in the same way as does pain perception. The behavioral part of this study indicates that placebo treatment has an effect on how subjects perceive unpleasant pictures. Furthermore, event-related fMRI demonstrated that the same modulatory network, including the rostral anterior cingulate cortex and the lateral orbitofrontal cortex, is involved in both emotional placebo and placebo analgesia. These effects were correlated with the reported placebo effect and were predicted by the amount of treatment expectation induced on a previous day. Thus, the placebo effect may be considered to be a general process of modulation induced by the subjects’ expectations.

ARTICLE UPDATE - The role of the amygdala in human fear: Automatic detection of threat

Arne Öhman

Psychoneuroendocrinology, in press

Behavioral data suggest that fear stimuli automatically activate fear and capture attention. This effect is likely to be mediated by a subcortical brain network centered on the amygdala. Consistent with this view, brain imaging studies show that masked facial stimuli activate the amygdala as do masked pictures of threatening animals such as snakes and spiders. When the stimulus conditions allow conscious processing, the amygdala response to feared stimuli is enhanced and a cortical network that includes the anterior cingulate cortex and the anterior insula is activated. However, the initial amygdala response to a fear-relevant but non-feared stimulus (e.g. pictures of spiders for a snake phobic) disappears with conscious processing and the cortical network is not recruited. Instead there is activation of the dorsolateral and orbitofrontal cortices that appears to inhibit the amygdala response. The data suggest that activation of the amygdala is mediated by a subcortical pathway, which passes through the superior colliculi and the pulvinar nucleus of the thalamus before accessing the amygdala, and which operates on low spatial frequency information.

ARTICLE UPDATE - The role of the human amygdala in the production of conditioned fear responses

David C. Knight, Hanh T. Nguyen and Peter A. Bandettini

NeuroImage, 26, 1193-1200

The amygdala plays a central role in the acquisition and expression of fear memories. Laboratory animal studies indicate that the amygdala both receives sensory information and produces learned behavioral and autonomic fear responses. However, prior functional imaging research in humans has largely focused on amygdala activity elicited by fearful stimuli, giving less attention to this region's role in the production of fear responses. In contrast, the present study used functional magnetic resonance imaging to investigate the amygdala's influence on the generation of conditional fear responses. Significant increases in amygdala activity were observed during the production of conditioned (learning-related), but not orienting, nonspecific, and unconditioned (nonlearning-related) skin conductance responses. Further, greater amygdala activity was demonstrated during conditioned response production than during conditioned stimulus presentation. These results suggest the amygdala not only responds to fearful stimuli, but also generates learning-related changes in human autonomic fear expression.

ARTICLE UPDATE - Hemodynamic responses of the amygdala, the orbitofrontal cortex and the visual cortex during a fear conditioning paradigm.

Tabbert K, Stark R, Kirsch P, Vaitl D

International Journal of Psychophysiology, 57, 15-23.

Functional magnetic resonance imaging (fMRI) studies consistently demonstrate an enhanced activation of the visual cortex in reaction to emotionally salient visual stimuli. This increase of activation is probably modulated by top-down processes, that are initiated in emotion processing structures, specifically the amygdala and the orbitofrontal cortex. In the present fMRI study, a differential fear conditioning paradigm was applied to investigate this assumed modulation. Hemodynamic responses towards a neutral visual stimulus (CS+) predicting an electrical stimulation (UCS) were compared with responses towards a neutral and unpaired stimulus (CS-). Thereby, particularly the time courses of neural responses were considered. Skin conductance measures were concurrently recorded. Our results show that the differentiation between CS+ and CS- within the amygdala and the extended visual cortex was accomplished during a late acquisition phase. In the orbitofrontal cortex the differentiation occurred at an earlier stage and was then sustained throughout acquisition. It is suggested that these altering activation patterns are reflecting different phases of learning, integrating the analyzed regions to varying degrees. Additionally, the results indicate that statistical analyses comprising a temporal variation of hemodynamic responses are more likely to detect amygdala activation.

ARTICLE UPDATE - Stimulus type and design influence hemodynamic responses towards visual disgust and fear elicitors.

Schafer A, Schienle A, Vaitl D.

International Journal of Psychophysiology, 57, 53-59.

The question to what extent emotion-related brain activation depends upon the presentation design (block design vs. event-related design) and the stimulus type (scene pictures vs. pictures with facial mimic) has hardly been addressed in previous functional magnetic resonance imaging (fMRI) research. In the present fMRI experiment, 40 right-handed subjects viewed pictures with fear-inducing and disgust-inducing content as well as facial expressions of fear and disgust. Pictures of neutral objects and neutral facial mimic were used as control stimuli. The pictures were presented in a block design for half of the subjects; the other half viewed the same stimuli as singular events in randomized sequence. The participants had been instructed to passively view the pictures. Disgust-evoking scenes provoked activation in the amygdala, the insula and the orbitofrontal cortex (OFC). This applied to the blocked as well as to the event-related design. Fear-relevant scenes were associated with activity in the insula, the OFC and the middle temporal gyri in the event-related design. The presentation in a block design only led to activation in the middle temporal gyri. Facial expressions of disgust and fear did not trigger significant activation neither in the blocked nor event-related design. This surprising outcome may be a result of context and task effects. The face stimuli which were presented together with the more complex scenes in a passive viewing paradigm possibly were not salient enough to trigger emotional processing.

ARTICLE UPDATE - The interaction of attention and emotion

John G. Taylor & Nickolaos F. Fragopanagos

Neural Networks, in press

We analyse emotions from the viewpoint of how emotion and attention interact in the brain. Much has been learnt about the brain structures involved in attention, especially in vision. In particular the manner in which attention functions as a high-level control system, able to make cognitive processing so effective, has been studied both at a global level by brain imaging (fMRI, PET, MEG and EEG), at a local single cell level in monkeys and lower animals, and computationally by a variety of models. The manner in which emotions impinge on this attention control system is not so well analysed, although numerous new results are now emerging from using the same tools. Here we use an engineering control approach to attention to model it in a global manner but with relatively sure local foundations at singe neuron level.

The manner in which emotional value (as coded in amygdale and orbito-frontal cortex) can interact with the attention control circuitry is analysed using results of various experimental paradigms. A general model of this interaction is first developed and tested against a list of paradigms, and then more detailed computations are performed using more specific features of the attention control system and the limbic value coding. These computations are completed by a simulation of the emotional attentional blink, a demanding paradigm for any model of attention alone, but made more so by the presence of emotional value codes for stimuli. We conclude the paper with a general discussion of further avenues of research.

ARTICLE UPDATE - Retrieving accurate and distorted memories: Neuroimaging evidence for effects of emotion

Elizabeth A. KensingeR & Daniel Schacter

NeuroImage, in press

While limbic activity is known to be associated with successful encoding of emotional information, it is less clear whether it is related to successful retrieval. The present fMRI study assessed the effects of emotion on the neural processes engaged during retrieval of accurate compared to distorted memories. Prior to the scan, participants (16 young adults) viewed names of neutral (e.g., frog) and emotional (e.g., snake) objects and formed a mental image of the object named. They were shown photos of half of the objects. During the fMRI scan, participants saw object names and indicated whether or not they had seen the corresponding photo. Memory distortions (misattributions) occurred when participants incorrectly indicated whether or not a photo had been studied. Activity in some regions (e.g., L anterior hippocampus) was related to accurate retrieval (correct attributions > misattributions) for emotional and neutral items. However, activity in other regions corresponded with accurate retrieval specifically for emotional items (e.g., in R amygdala/periamygdaloid cortex and L orbitofrontal cortex) or for neutral items (e.g., in lateral inferior prefrontal cortex and R posterior hippocampus). Results indicate that emotional salience modulates the processes engaged during accurate retrieval and that activity in limbic regions corresponds with accurate memory assignment for emotional items. To our knowledge, this study is the first to demonstrate a link between limbic engagement at retrieval and accurate memory attribution.

ARTICLE UPDATE - The neural bases of amusement and sadness: A comparison of block contrast and subject-specific emotion intensity regression approache

Goldin PR, Hutcherson CA, Ochsner KN, Glover GH, Gabrieli JD, Gross JJ.

Neuroimage, in press


Neuroimaging studies have made substantial progress in elucidating the neural bases of emotion. However, few studies to date have directly addressed the subject-specific, time-varying nature of emotional responding. In the present study, we employed functional magnetic resonance imaging to examine the neural bases of two common emotions-amusement and sadness-using both (a) a stimulus-based block contrast approach and (b) a subject-specific regression analysis using continuous ratings of emotional intensity. Thirteen women viewed a set of nine 2-min amusing, sad, or neutral film clips two times. During the first viewing, participants watched the film stimuli. During the second viewing, they made continuous ratings of the intensity of their own amusement and sadness during the first film viewing. For sad films, both block contrast and subject-specific regression approaches resulted in activations in medial prefrontal cortex, inferior frontal gyrus, superior temporal gyrus, precuneus, lingual gyrus, amygdala, and thalamus. For amusing films, the subject-specific regression analysis demonstrated significant activations not detected by the block contrast in medial, inferior frontal gyrus, dorsolateral prefrontal cortex, posterior cingulate, temporal lobes, hippocampus, thalamus, and caudate. These results suggest a relationship between emotion-specific temporal dynamics and the sensitivity of different data analytic methods for identifying emotion-related neural responses. These findings shed light on the neural bases of amusement and sadness, and highlight the value of using emotional film stimuli and subject-specific continuous emotion ratings to characterize the dynamic, time-varying components of emotional responses.

ARTICLE UPDATE - A direct intracranial record of emotions evoked by subliminal words

Lionel Naccache, Raphaël Gaillard, Claude Adam, Dominique Hasboun, Stéphane Clémenceau, Michel Baulac, Stanislas Dehaene, and Laurent Cohen

PNAS, 102, 7713-7717.

A classical but still open issue in cognitive psychology concerns the depth of subliminal processing. Can the meaning of undetected words be accessed in the absence of consciousness? Subliminal priming experiments in normal subjects have revealed only small effects whose interpretation remains controversial. Here, we provide a direct demonstration of semantic access for unseen masked words. In three epileptic patients with intracranial electrodes, we recorded brain potentials from the amygdala, a neural structure that responds to fearful or threatening stimuli presented in various modalities, including written words. We show that the subliminal presentation of emotional words modulates the activity of the amygdala at a long latency (>800 ms). Our result indicates that subliminal words can trigger long-lasting cerebral processes, including semantic access to emotional valence.

ARTICLE UPDATE - Binocular rivalry between emotional and neutral stimuli: A validation using fear conditioning and EEG

Georg W. Alpers, Mirjana Ruhleder, Nora Walz, Andreas Mühlberger and Paul Pauli

International Journal of Psychophysiology, in press

When two incompatible pictures are projected to the two eyes, they compete for perceptual dominance. Previous research has claimed that meaningful and emotionally valenced pictures predominate over neutral pictures in this rivalry. This may be interpreted as evidence for preferential processing of emotionally significant stimuli in the visual system but it is difficult to dismiss that the physical characteristics of the different pictures or response biases influenced the results of these studies. Thus, we set out to examine the influence of emotion using methods eliminating the influence of physical characteristics and minimizing response biases. We used simple visual patterns and induced emotional valence by fear conditioning. In Experiment 1 the aversive CS+ predominated over the CS−. In Experiment 2 we extended previous findings by showing that participants' self-reported perception is validated by corresponding steady-state visually evoked potentials in the EEG in the context of such a conditioning experiment. This was accomplished by frequency coding the rivalling stimuli with a stimulus specific pattern reversal and extracting the corresponding frequency from the occipital lobe EEG. Taken together, these studies provide further evidence that picture valence can influence perception in binocular rivalry. This is discussed in terms of subcortical mechanisms supporting the efficient processing of threatening information.

ARTICLE UPDATE - Functional connectivity with anterior cingulate and orbitofrontal cortices during decision-making

M.X Cohen, A.S. Heller and C. Ranganath

Cognitive Brain Research, 23, 61-70

Recent neuroscience research is beginning to discover the brain regions involved in decision-making under uncertainty, but little is known about whether or how these regions functionally interact with each other. Here, we used event-related functional magnetic resonance imaging to examine both changes in overall activity and changes in functional connectivity during risk-taking. Results showed that choosing high-risk over low-risk decisions was associated with increased activity in both anterior cingulate and orbitofrontal cortices. Connectivity analyses revealed that largely distinct, but somewhat overlapping, cortical and subcortical regions exhibited significant functional connectivity with anterior cingulate and orbitofrontal cortices. Additionally, connectivity with the anterior cingulate in some regions, including the orbitofrontal cortex and nucleus accumbens, was modulated by the decision participants chose. These findings (1) elucidate large networks of brain regions that are functionally connected with both anterior cingulate and orbitofrontal cortices during decision-making and (2) demonstrate that the roles of orbitofrontal and anterior cingulate cortices can be functionally differentiated by examining patterns of connectivity.

ARTICLE UPDATE - The cognitive control of emotion

Kevin N. Ochsner and James J. Gross

Trends in Cognitive Sciences, 9, 242-249

The capacity to control emotion is important for human adaptation. Questions about the neural bases of emotion regulation have recently taken on new importance, as functional imaging studies in humans have permitted direct investigation of control strategies that draw upon higher cognitive processes difficult to study in nonhumans. Such studies have examined (1) controlling attention to, and (2) cognitively changing the meaning of, emotionally evocative stimuli. These two forms of emotion regulation depend upon interactions between prefrontal and cingulate control systems and cortical and subcortical emotion-generative systems. Taken together, the results suggest a functional architecture for the cognitive control of emotion that dovetails with findings from other human and nonhuman research on emotion.

ARTICLE UPDATE - Same or different? Neural correlates of happy and sad mood in healthy males

Ute Habel, Martina Klein, Thilo Kellermann, N. Jon Shah and Frank Schneider

NeuroImage, 26, 206-214

Emotional experience in healthy men has been shown to rely on a brain network including subcortical as well as cortical areas in a complex interaction, which may be substantially influenced by many internal personal and external factors such as individuality, gender, stimulus material and task instructions. The divergent results may be interpreted by taking these considerations into account. Hence, many aspects remain to be clarified in characterizing the neural correlates underlying the subjective experience of emotion. One unresolved question refers to the influence of emotion quality on the cerebral substrates. Hence, 26 male healthy subjects were investigated with functional magnetic resonance imaging during standardized sad and happy mood induction as well as a cognitive control task to explore brain responses differentially involved in positive and negative emotional experience. Sad and happy mood in contrast to the control task produced similarly significant activations in the amygdala–hippocampal area extending into the parahippocampal gyrus as well as in the prefrontal and temporal cortex, the anterior cingulate, and the precuneus. Significant valence differences emerged when comparing both tasks directly. More activation has been demonstrated in the ventrolateral prefrontal cortex (VLPFC), the anterior cingulate cortex (ACC), the transverse temporal gyrus, and the superior temporal gyrus during sadness. Happiness, on the other hand, produced stronger activations in the dorsolateral prefrontal cortex (DLPFC), the cingulate gyrus, the inferior temporal gyrus, and the cerebellum. Hence, negative and positive moods reveal distinct cortical activation foci within a common neural network, probably making the difference between qualitatively different emotional feelings.